Alena [Aztekoraphidia) Horstaspoecki Nov. Spec. - a New Snakefly from Mexico (Raphidioptera, Raphidiidae)1

Denisia 13 | 17.09.2004 | 129-134

Alena [Aztekoraphidia) horstaspoecki nov. spec. - a new snakefly from Mexico (Raphidioptera, Raphidiidae)1

U. ASPÖCK Et A. CONTRERAS-RAMOS

Abstract: Alena {Aztekoraphidia) horstaspoecki nov. spec, a new species of snakefly, from Hidalgo state, central Mex- ico, is described and illustrated. With this discovery the number of snakefly species recorded from Mexico increases to 14- Morphological criteria of the hypovalva reveal diagnostic characters for the differentiation from all other species of Alena, and support arguments for the hypothesis of a hypovalva-paramere-complex.

Resumen: Una especie nueva de rafidiöptero, Alena (Aztekoraphidia) horstaspoecki nov. spec, es diagnosticada, de- scrita e ilustrada con ejemplares del estado de Hidalgo, en el centro de Mexico. Esta es la d£cimocuarta especie de rafidiöptero registrada en Mexico. Criterios morfolögicos de la hipovalva revelan caracteres diagnösticos para la se- paration de todas las demäs especies de Alena, apoyando ademäs la hipötesis de un complejo hipovalva-paramero.

Key words: Raphidioptera, Alena {Aztekoraphidia), new species, Mexico.

Introduction Alena [Aztekoraphidia) horstaspoecki nov. spec. The Raphidioptera is a small order which comprises two families, the Raphidiidae with 186 described valid Derivatio nominis: The name of this new species is a species, and the Inocelliidae with 21. Raphidioptera to- grateful homage to Horst Aspöck, Vienna, Austria, for his extensive contribution to neuropterology, on the oc- gether with its adelphotaxon Megaloptera + Neuroptera casion of his 65 th birthday. constitute the Neuropterida, a superorder at the base of the Holometabola. The Raphidioptera is distributed Material studied: Holotype, C: Mexico, Hidalgo, Huasca, Ran- throughout the Holarctic region, except for the northern cho Sta. Elena, Manantial de las Vigas, 20°07'53,4"N/ and eastern parts of North America, the southernmost 98°31'38,5"W, 2500 m, 26.I1I.-9.IV.2OO3, Menchaca, Contr- : records being from high altitudes at the Mexican- eras; Malaise 1; Paratypes: 1 9 Same data as holotype, but 9.- Guatemalan border, northern Africa, northern India, 23.1V.; 1 9: Same locality as holotype, 26.11.03-26.111.03, Myanmar, northern Thailand and Taiwan (H. ASPÖCK, Malaise 2; (cr holotype and one 9 paratype in coll. Centro de Investigaciones Biologicas, UAEH, Pachuca, Mexico, one 9 U. ASPÖCK & RAUSCH 1991). In Mexico, hitherto 13 paratype in coll. Naturhistorisches Museum Wien). species have been recorded, 10 Raphidiidae, and three Morphology (Fig. 1-11): A small tender species, Inocelliidae. All available information on Mexican length of forewing of the O" 8 mm, of the 9 8,2-9 mm. snake flies has been summarized in U. ASPÖCK &. H. As- Head elongate rectangular, black with brownish pattern, PÖCK (1996). The discovery of a new species in this part with a fine sculpture, clypeus brownish, with a group of of the earth is not at all surprising, nevertheless it is of bristles; labrum brownish with yellow margins. Anten- great significance for several reasons: 1) for a relict taxon nae: scapus brown, pedicellus and basal flagellomeres yel- comprising altogether 207 extant described species the lowish, otherwise brown. Pronotum: in front and lateral- discovery of species number 208 is a spectacular event, 2) ly with broad yellow margins, anterior part brownish, the new species is from a region of Mexico where so far longer caudal part dark brown, almost black, with ochre no raphidiopterans have been reported, 3) it documents pattern. Legs: first coxae basally brown, distally yellow, once more the importance of the Mexican region as an evolutionary centre of these living fossils, and 4) a re- 1 Dedicated to Horst Aspöck on the occasion of his 65th birthday, grate- markable anatomical character of the hypovalva induces fully and in all the excitement of 40 years fascinating collaboration on a new interpretation of this structure. Neuropterida, and of continuous stimulating scientific discussions. Ul- rike Aspöck. Dedicated to Prof. Horst Aspöck, gladly and as a great honour, on the occasion of his 65th birthday. Atilano Contreras-Ramos.

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Fig. 1: Alena {Aztekoraphidia) horstospoecki nov. spec, holotype, er, head Fig. 2: Alena (Aztekorophidia) horstaspoecki nov. spec, holotype, er, and thorax, lateral. abdomen and wings, ventral.

Fig. 3: Alena (Aztekoraphidia) horstospoecki nov. spec, holotype, er, genital Fig. 4: Alena [Aztekoraphidia) horstaspoecki nov. spec, holotype, er, genital segments, ventral. segments, lateral.

Fig. 5: -• r'\; Aztekoraphidia) horstaspoecki nov. spec, paratype, Fig. 6: Alena [£.••••• . •.:;. "idia) horstaspa:- spec, paratype, Q, abdomen and wings, ventral. genital segments, ventral.

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0,5 mm 0,5 mm Fig. 7: Alena [Aztekorophidia) horstospoecki nov. spec, holotype, c, genital Fig. 8: Aleno [Aztekoraphidia) horstaspoecki nov. spec, holotype, cr, genital segments, lateral, e = ectoproct, gc9 = 9th gonocoxite, h = 9th segments, dorsal. Abbreviations as in fig. 7. gonapophysis (hypovalva), h + p? = amalgamation of hypovalva and parameres, hi = hypandrium internum, p ? = 10th gonocoxite + 10th gonapophysis ? (paramere?), st = stylus.

gphS

Fig. 10: Alena (Aztekorophidia) horstospoecki nov. spec, paratypus, 9, genital segments, lateral, ab = atrium bursae, abh = „horn" of atrium Fig. 9: Alena {Aztekoraphidia) horstaspoecki nov. spec, holotype, cr, genital bursae, e = ectoproct, gc9 = 9th gonocoxites, gph8 = 8th gonapophyses, gr segments, ventral. Abbreviations as in fig. 7. = glandulae receptaculi, rs = receptaculum seminis, sb = sacculus bursae. second and third coxae brown; tarsalia 3-5 brown, legs otherwise yellowish. Wings: membrane hyaline, vena- 0,5 mm tion predominantly brown, basal half of C, basal R and Cu yellowish; basal part of Ma in hind wing as a longitu- gc9 dinal vein. Pterostigma yellow, above the distal half of the first pterostigmal cell, with one vein crossing, and with apical vein. Abdomen: tergites speckled brownish; sternites anteriorly black, with two yellow spots in female, but two large yellow windows in male; caudally yellow, medially with a brownish deltoid patch; pleural region brownish-yellowish.

cr genital segments (Fig. 3-4, 7-9): Sternite of segment 8 only slightly shorter than tergite; tergite 9 trapez- Fig. 11: Alena (Aztekoraphidia) horstospoecki nov. spec, paratype, 9, iform; gonocoxites 9 forming huge plates with a blunt genital segments, ventral. short, yellowish-whitish, only proximally and basally brownish apex; styli slim; sternite 8 rather indistinct; hypovalva paired, with broad basal rods, dorsally curved

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spiraculum 7 free; tergite 8 inconspicuous, not reaching ventral part of the segment. Atrium bursae ventrally with paired „horns", joined to a complex together with the receptaculum seminis, which is, however, still discernible; sacculus bursae and a pair of glandulae receptaculi skinny and hose-shaped, apically club-like enlarged. Differentiation: According to the morphology of the ninth gonocoxites of the male A. horstaspoecki nov. spec, belongs to the species-group of the subgenus Azteko- raphidia, which is constituted by A. caudata (NAVAS 1914), A. minuta (BANKS 1903), A. australis (BANKS 1895), A. tenochtitlana U. ASPÖCK & H. ASPÖCK 1978, A. mfundihdata U. ASPOCK & H. ASPÖCK & RAUSCH 1994, and A. schremmeri U. ASPÖCK & H. ASPOCK & RAUSCH 1994, without being closely related to any of them. It may be distinguished from all other species of the subgenus Aztekoraphidia, but also of the genus Alena, by the huge and heavily sclerotized lateral plates of the hypovalva, by the blunt yellowish-whitish apex of the ninth gonocoxites in the

Habitat (Fig. 12, 13): The type locality consists of two sites, one adjacent to a first order stream close to its origin (Malaise 1), and a second one at about 20 meters from the stream into the forest in a fairly open area (Malaise 2). The vegetation consists of a pristine oak- pine forest at the base of a mountainous area. Both Malaise traps were maintained operating continuously for one year. However, the three snakefly specimens on which this paper is based were the only ones collected throughout the sampling period.

Almost nothing is known on the biology of the new species, its discovery in a pine-oak forest may, however, lead to the conclusion that the larvae are corticolous, as are all Alena species whose larvae are known (H. ASPÖCK 2002). Distribution (Map 1): So far A. horstaspoecki is known from the type locality only, and most probably it is another endemic species of the genus, which now com-

Fig. 12 and ij .. ality of Alena(Aztekorophidia) prises seven species endemic to Mexico, and, in addition, horstaspoecki nov. spec. A. disancta (BANKS 1911) and A. minuta (BANKS 1903), the only species with a larger distribution area north of apices, and a pair of strongly sclerotized lateral plates; ec- Mexico. toproct with distinct trichobothria; hypandrium internum inconspicuous.

9 genital segments (Fig. 5, 6, 10, 11): Caudal mar- gin of sternite 7 convex with weak median incision,

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Indianocellia r-,•. . Nagha maridonaiis Agula bnofcx AgiAaamaudl Awns oitinctB AJana Mundbütti

Fig. 14: Records of Raphidoptera in Mexico. For exact data of localities see in H. ASPOCK, ASPÖCK U. ft RAUSCH (1991), U. ASPÖCK (1974, 1975, 1988), U. ASPÖCK ft H. ASPOCK (1970, 1976, 1978, 1996, and U. ASPÖCK, ASPÖCK H. ft RAUSCH (1992, 1994a, 1994b).

Discussion RAUSCH 1991, Fig. 1036, 1037), Italoraphidia solariaw (NAVÄS 1928) (see H. ASPÖCK, U. ASPÖCK &. RAUSCH The anatomy of the raphidiopteran hypovalva, which 1991, Fig. 1069,1070). The homologisation of these scle- has been interpreted as the gonapophyses of segment 9 rites (U. ASPÖCK 2002) as parameres (tenth gonocoxites (U. ASPÖCK 2002), deserves some further considerations. + tenth gonapohyses) or, alternatively, as the gonarcus In the family Raphidiidae the hypovalva is a paired or un- (eleventh gonocoxites), has always been problematic. On paired sclerite, closing the male terminalia ventrally. The the other hand, there are similar structures, accompany- family Inocelliidae lacks a comparable hypovalva, but the ing the hypovalva laterally, partly externally and more ninth gonapophyses are still to be found as the so called basally, which have been identified as parameres, e.g. in pseudostyli in some species. The hypovalva may be of the Palaearctic Raphidia (R.) ophiopsis LINNAEUS 1758, or very different shape, and at any rate it is of enormous tax- in the Nearctic Aguila {Glavia) modesta CARPENTER 1936 onomic and systematic importance. Based on considera- (see H. ASPÖCK, U. ASPÖCK & RAUSCH 1991, Fig. 771- tions turned over already long ago the hitherto unpub- 810 and Fig. 1794-1845) respectively. lished hypothesis of the composed nature of the hypovalva of certain Raphidiidae is corroborated by male genital Under these aspects the study of the new species of- characters of the new species. This is particularly obvious fers the following hypotheses: in species of the Nearctic subgenus Aztekoraphidia U. 1) The apical part of the hypovalva, comprising the ASPÖCK & H. ASPÖCK 1970 of the genus Alena NAVÄS lateral plates and the curved hooks are interpreted as the 1916, but probably also of the Mediterranean genus His- amalgamated parameres (gonocoxites + gonapophyses of panoraphidia H. ASPÖCK & U. ASPÖCK 1968. The basal segment 10). parts and the apical parts of the paired rods of the hypovalva show traces of a disruption, in a way or the other, ei- 2) In other species of Aztekoraphidia - although not ther by skinny parts, or by special structures, e.g. Alena evident at a first glance, as the lateral plates are missing (Aztekoraphidia) caudata, A. minuta, A. australis (BANKS - the hypovalva too seems to be of composed nature, and 1895), or A. tenochtidana (U. ASPOCK & H. ASPÖCK the apices represent the amalgamated parameres. 1978). In A. horstaspoecki the apical hooks and the later- 2) If the hypothesis of amalgamation of parameres al plates of the hypovalva appear as if they were amalga- and hypovalva holds, identical phenomena in Palaearc- mated structures, and resemble certain internal sclerites tic species could be of phylogenetic relevance, if they are known from Palaearctic species: e.g. Adantoraphidia ma- synapomorphic. The genera Harraphidia STEINMANN culkolks (STEPHENS 1836) (see H. ASPÖCK, U. ASPÖCK & 1963 and Hispanoraphidia H. ASPOCK & U. ASPÖCK 1968

have to be considered under this aspect, as both lack ASPÖCK U. & H. ASPÖCK (1970): Untersuchungen über die Raphi- parameres, and the hypovalva might include the para- diopteren Mexikos (Insecta, Neuropteroidea). — Polskie Pis- mo ent. 40: 705-725. meres in these (and perhaps additional) taxa. Moreover, this could be also of biogeographic relevance for an un- ASPOCK U. & H. ASPOCK (1976): Zur Taxonomie und Systematik von Raphidia lAliaberaphidia) australis BANKS (Ins., Neun, Neu- derstanding of the origin of the Nearctic Raphidiidae. roptera). — Ent. Ben, Amst. 36: 137-139.

ASPOCK U. & H. ASPOCK (1978): Raphidia (Aliaberaphidia) tenoch- Acknowledgements titlana n.sp. - eine neue Raphidiiden-Spezies aus Baja Cali- fornia (Mexiko) (Neuropteroida, Raphidioptera). — Z. Arb- Financial support for field work by the junior author from SEP- Gem. öst. Ent. 30: 21-24. PROMEP (Mexico), through the project Sistemätica de Tulipi- ASPOCK U. & H. ASPOCK (1996): Raphidioptera. — In: BOUSOUETS dae (Diptera) y Neuroptera del Estado de Hidalgo, as well as J.E.L., AIDRETE A.N.G. & E.G. SORIANO (eds), Biodiversidad, Ta- permission and support for collecting in Rancho Santa Elena xonomia y Biogeografia de Artröpodos de Mexico: Hacia from Cecilia Chävez-Peön and Roberto Campuzano, are grate- una sintesis de su conocimiento 19: 277-286, D.R. Universi- fully acknowledged. dad nacional Autön. Mexico. Cordial thanks to Franziska Änderte, University Vienna, for ASPOCK U., ASPOCK H. & H. RAUSCH (1992): Rezente Südgrenzen der graphical assistance with the drawings of male and female gen- Ordnung Raphidioptera in Amerika (Insecta: Neuropteroi- italia, and the distribution map, and to Dr. Martin Lödl, dea). — Entomol. Gener. 17: 169-184. Naturhistorisches Museum Wien, for photographing the types. ASPOCK U„ ASPOCK H. S H. RAUSCH (1994a): Neue Arten der Familie Finally we want to thank Robert Seiler, Naturhistorisches Mu- Raphidiidae aus Mexiko und Nachweis einer Spermatopho- seum Wien, for his continuous help in all computer matters. re in der Ordnung Raphidioptera (Insecta: Neuropteroidea). —Entomol. Gener. 18: 145-163. Zusammenfassung ASPOCK U., ASPOCK H. & H. RAUSCH (1994b): Alena (Mexicoraphidia) Alena {Aztekoraphidia) horstaspoecki nov. spec, eine neue Ka- americana (CARPENTER, 1958): Taxonomie, Systematik, Ökolo- melhalsfliege aus Hidalgo, Zentralmexiko, wird beschrieben gie und Chorologie (Neuropteroidea: Raphidioptera: Raphi- diidae).—Z. ArbGem. öst. Ent. 46: 131-139. und abgebildet. Mit dieser Neuentdeckung erhöht sich die Zahl der bisher aus Mexiko nachgewiesenen Spezies der Ordnung Raphidioptera auf 14- Morphologische Kriterien der Hypovalva bieten nicht nur diagnostische Merkmale zur Abgrenzung gegenüber allen anderen Akna-Arten, sondern liefern auch Ar- gumente zur Hypothese eines Hypovalva-Parameren-Komple-

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